Redescription of the subgenus Parancistrum Verhoeff

Zootaxa 3734 (5): 501–520 www.mapress.com /zootaxa / Copyright © 2013 Magnolia Press

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http://dx.doi.org/10.11646/zootaxa.3734.5.1 http://zoobank.org/urn:lsid:zoobank.org:pub:2C4411BF-233F-48F2-BA6F-423A9F8EE8B1

Redescription of the subgenus Parancistrum Verhoeff, 1943, an eastern Mediterranean lineage of the millipede genus Megaphyllum Verhoeff, 1894 (Diplopoda: Julida: Julidae: Brachyiulini) BOYAN VAGALINSKI1,2, ESZTER LAZÁNYI3 & SERGEI GOLOVATCH4 1

Faculty of Biology, Sofia University, 8 Dragan Tsankov Blvd., 1164 Sofia, Bulgaria. E-mail: [email protected] Institute of Biodiversity and Ecosystem Research, Bulgarian Academy of Sciences, 2 Yurii Gagarin Street, 1113, Sofia, Bulgaria 3 Department of Zoology, Hungarian Natural History Museum, Budapest, Baross u. 13, H-1088, Hungary. E-mail: [email protected] 4 Institute for Problems of Ecology & Evolution, Russian Academy of Sciences, Leninsky prospect 33, Moscow 117051, Russia. E-mail: [email protected] 2

Abstract The subgenus Parancistrum Verhoeff, 1943 is redefined on the basis of the examination of type and non-type material. The subgenus is currently settled to comprise 6 species, of which Megaphyllum arcuatum sp. n. is described as new, while M. adanense (Verhoeff, 1943) syn. n. and M. palaestinum (Jawłowski, 1931) syn. n. are established as junior subjective synonyms of M. curvifolii (Verhoeff, 1898) and M. genezarethanum (Verhoeff, 1923) comb. nov., respectively. A neotype is designed for Chromatoiulus tenenbaumi Jawłowski, 1931 under ICZN Article 75.1. The distribution pattern of the studied group is discussed, and some preliminary phylogenetic ideas about Megaphyllum and the tribe Brachyiulini are presented. Key words: millipede, Brachyiulus, new species, new synonymies, neotype, distribution

Introduction The mostly Euro-Ponto-Mediterranean genus Megaphyllum Verhoeff, 1894 is one of the most diverse and taxonomically complex genera in the family Julidae (Akkari et al. 2011; Lazányi et al. 2012). Its identity has been entangled with numerous problems across its history. Numerous assigned species have repeatedly been referred to under different generic names, such as Julus Linnaeus, 1758, Brachyiulus Berlése, 1884 and/or Chromatoiulus Verhoeff, 1894; mostly as a result of Verhoeff’s inconsistent views which often contradicted the basic rules of the ICZN (see also Pocock 1895). It was Hoffman (1980) who finally resolved the confusion by giving clear arguments for re-establishing the name “Megaphyllum” (see also Lazányi & Korsós 2011). However, a large number of highly varied species were assigned to the genus, mostly due to the lack of a diagnosis and the very general original description. The attempts at intra-generic grouping have even increased the ambiguity, resulting in a real jungle of numerous, mostly monotypic, sometimes overlapping or even invalidly proposed subgenera. Parancistrum Verhoeff, 1943 is one of these problematic subgenera. It was diagnosed on the basis of a species which is actually a junior synonym of M. curvifolii (Verhoeff, 1898), and has since remained monotypic despite the presence of several other, obviously similar species already known at that time. Verhoeff (1898) described Megaphyllum curvifolii and M. turcicum from “Cilicien” [near Tarsus], and M. serratum from Pernata, both in Turkey. Jawłowski (1931) described two new Megaphyllum species: M. palaestinum and M. tenenbaumi, both from today’s Israel, without referring to the above mentioned Verhoeff’s paper. Meanwhile, Verhoeff (1923) had already described a new species of the genus, namely M. genezarethanum, from the same area, based on a single female. This was noted by Jawłowski (1931), who mentioned a possible synonymy of M. genezarethanum with one of his two new species. Attems (1940), in his revisionary work on Julidae, placed M. curvifolii, M. serratum, M. palaestinum and M. tenenbaumi, as well as a number of other species, in a newly described subgenus of Megaphyllum (Chromatoiulus auct.), namely, Phauloiulus Attems, 1940. Accepted by W. Shear: 15 Oct. 2013; published: 8 Nov. 2013

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However, this subgenus can hardly be treated as a natural group (see comment under the description of Parancistrum). Verhoeff (1943), in his work on Turkish millipedes, described M. adanense and proposed the newly described subgenus Parancistrum Verhoeff, 1943 to encompass that species alone. In the same paper, Verhoeff failed to refer to any of the previous publications, including his own ones which dealt with Megaphyllum from the Near East region. As the next step in our ongoing Megaphyllum revision, following a revision of the subgenus Megaphyllum sensu stricto (Lazányi & Vagalinski, accepted), we here redefine the subgenus Parancistrum, a generally overlooked, but distinct group of Megaphyllum.

Material and methods All studied specimens are preserved in 70% ethanol and are deposited in the following museums: Museum für Naturkunde, Berlin (MNKB); Muséum d’histoire naturelle, Geneva (MHNG); Naturhistorisches Museum, Wien (NHMW); Natural History Museum of Denmark, Zoological Museum, University of Copenhagen (ZMUC); Zoological Museum, University of Moscow (ZMUM); Zoologische Staatssammlung, München (ZSM); Zoological Museum, Tel Aviv University (ZMTA); National Museum of Natural History, Sofia (NMNHS), and Hungarian Natural History Museum (HNHM). Body measurements were taken as follows: body length was measured at ozopore level, body height at midbody ring; antenno- and podomeres were measured according to Enghoff (1992). Developmental stadium data and the number of body rings are presented in the following formula: stadium number, number of podous rings+number of apodous rings+T, where T is the telson (after Enghoff et al. 1993). Gonopods, vulvae and penes were dissected and mounted in temporary glycerin or permanent Eukitt slide mounts for examination and drawing. The internal structures of the vulvae were clarified using a 10% NaOH solution. Drawings were made using a MБC-9 (SU) or Leica M125 (HNHM) and Leica MZ125 (NHMW) compound microscopes. Scanning electron micrographs were taken with JEOL JSM-5510 (SU) or JEOL JSM6335F (ZMUC) scanning electron microscopes. Photographs were obtained using Olympus stereomicroscopes. The terminology of the various gonopodal and vulval parts is as follows: posterior opisthomere process (pp), anterior opisthomere process (ap), solenomere (s), anterior solenomere protrusion (asp), posterior solenomere process (psp), seti- or spine-like filaments (sf), flagellum (fl), flagellum channel (fc), promere (P), median ridge (r), median groove (g), operculum (op), operculum’s lamellar protrusions (opl), bursal lamellar protrusions (bl), opening (o), lateral hump (lh), central hump (ch), mesal hump (mh), posterior ampulla (pa), apodematic tube (at) and connecting tube (ct).

Taxonomic part Genus Megaphyllum Verhoeff, 1894 Subgenus Parancistrum Verhoeff, 1943 Parancistrum Verhoeff, 1943: 226. Type species: Brachyiulus adanensis Verhoeff, 1943 (syn. n. of Megaphyllum curvifolii (Verhoeff, 1898)), by monotypy.

Diagnosis. A subgenus of Megaphyllum differing from other subgenera by the following combination of characters: Opisthomere with a large, flattened, posterior process (pp), a well-developed, slender, club- or spineshaped anterior process (ap), and a rather simple solenomere, with a prominent anterior protrusion (asp). Vulva cylindrical, with a small, entirely apically positioned opening (o); operculum (op) shorter than to subequal to bursa. Subanal scale in males triangular, pointed. Description. Medium- to large-sized animals, 25–45 mm long and 1.6–3.2 mm high; females usually slightly longer and considerably thicker than males. Body colour pattern usually marked by two lateral and one mid-dorsal dark stripe against a brownish background. Head with 2 frontal, 4 supralabral and 18–24 labral setae. Antennomeres 3, 4 and 5 subequal in length.

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Gnathochilarium typically julid, with 4–5 setae on each lingual plate; promentum rather large, 1/4–2/7 of total length of gnathochilarium. Males with expanded mandibular stipites, each protruding ventro-anteriad into a rounded lobe. Collum smooth, either with several short scratches at posterolateral corner or with one long stria running along collum’s lateral margin. Body rings not vaulted; male’s first 3–4 body rings somewhat flattened dorsoventrally, this being more pronounced in certain species. Prozona finely scratched, metazona deeply and densely striated. Male pleurotergum 7 with large shovel-shaped or smaller lobes protruding mesad at border between pro- and metazona. Pre-anal process (= epiproct) long and pointed, sometimes with a hyaline tip turned down, in females usually shorter and stouter. Anal valves (= paraprocts) densely pilose. Subanal scale (= hypoproct) triangular, pointed or faintly blunt, in females sometimes narrowly rounded. Male leg-pair 1 forming parallel hooks, without tarsal remnants; leg-pair 2 unmodified, with two adhesive pads, one each on postfemora and tibiae; following walking legs with three pads, one each on femora, postfemora and tibiae. Tarsi of mid-body male walking legs shorter than or subequal to tibiae. Legs in females unmodified except for first two leg-pairs being slightly enlarged when vulvae fully developed. Penis lying deep behind coxae 2, stout, somewhat longer than broad, with well-developed apical lobes slightly turned caudad. Gonopods more or less slender, in situ considerably protruding outside gonopod sinus. Promere (P) simple, elongated, leaf-like, with a rather deep median groove (g) on its posterior side and a well-developed median ridge (r). Flagellum (fl) from subequal to, to considerably longer than promere, thin; several spini- or setiform filaments (sf) along opisthomere’s flagellum channel (fc). Opisthomere somewhat longer than to subequal to promere, with a massive, flattened, posterior process (pp), a slender, smooth or serrated, spini- or claviform, anterior process (ap), and a rather simple solenomere (s) turned caudad and bearing a marked anterior protrusion (asp), with flagellum channel (fc) opening behind it. Vulvae robust, markedly cylindrical, with a small, slit-like, apically positioned opening (o); bursa (b) and operculum (op) subequal in height, with well-developed, apical, lamellar protrusions (bl and opl, respectively); setation particularly dense; receptaculum seminis double, consisting of an ovoid or subspherical posterior ampulla (pa) and a wide apodematic tube (at), the former attached to the latter through a long, folded, often spirally twisted connecting tube (ct). Distribution. Turkey, Israel Included species. Megaphyllum curvifolii (Verhoeff, 1898) (type species), M. genezarethanum (Verhoeff, 1923), M. serratum (Verhoeff, 1898), M. tenenbaumi (Jawłowski 1931), M. turcicum (Verhoeff, 1898) and M. arcuatum sp. n. Comment. Attems (1940) assigned the species M. curvifolii, M. genezarethanum (referred to as Chromatoiulus palaestinus), M. serratum, M. tenenbaumi and M. turcicum to the subgenus Phauloiulus Attems, 1940 he then described. However, by choosing M. lictor Attems, 1904 as the type species, a species actually belonging to Byzantorhopalum Verhoeff, 1932 (Vagalinski & Lazányi, in prep.), as well as by adding therein (in the same paper) also M. crassum (Attems, 1928), M. glossulifer (Schubart, 1934), M. karschi (Verhoeff, 1901), M. montivagum (Verhoeff, 1901), M. leucadium (Attems, 1929) (referred to as Chromatoiulus leucasius, a misprint), M. pentheri (Attems, 1905), M. seditiosum (Attems, 1940), M. tetricum (Attems, 1932) and M. vicinum (Verhoeff, 1903), Attems instantly made his Phauloiulus a highly heterogeneous assemblage. Moreover, M. curvifolii appears to be a senior subjective synonym of M. adanense, i.e. the type, and so far only constituent, species of Parancistrum. The decision to revive the hitherto monotypic Parancistrum is prompted by the presence of several similar species among Phauloiulus which, together with the synonymized M. adanense, form a natural group that shows certain similarities with Megaphyllum s. str., but is at the same time clearly discernible.

Megaphyllum (Parancistrum) curvifolii (Verhoeff, 1898) Figs 1–9 Brachyiulus curvifolii Verhoeff, 1898: 300–301, fig. 13. Brachyiulus curvifolii: Verhoeff 1923: 131. Chromatoiulus curvifolii: Attems 1927: 224. REDESCRIPTION OF SUBGENUS PARANCISTRUM

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Chromatoiulus (Phauloiulus) curvifolii: Attems 1940: 302. Brachyiulus adanensis Verhoeff, 1943: 231–232, figs 16–19, syn. n. Megaphyllum adanense: Enghoff 2006: 182. Megaphyllum curvifolii: Enghoff 2006: 183.

Material examined. MNKB: Brachyiulus curvifolii, ZMB 4379, 1 ♂, 1 ♀ syntypes, Verh.-slide 1199: gonopods, 1st, 2nd leg-pairs type slide preparation, Turkey, “Cilicien” [south-central Turkey], M. Holtz leg. NHMW: Brachyiulus (Chromatoiulus) curvifolii, Inv. No. 3103, 1 ♂, gonopods as slide preparation, 1 ♀, probably syntypes, Turkey, Cilicien. ZSM: M. adanense, 1 ♂, questioned type, Reg. Nr. A20060652, Asia, Turkey, Adana; Reg. Nr. A20041622 slide preparation: gonopods, gnathochilarium, antenna, 1st, 2nd, 3rd leg-pairs and some other legs; Reg. Nr. A20041623 slide preparation: same body parts. MHNG: Chromatoiulus curvifolii, 1 ♂, 1 subad. 1 ♀, Asie Mineure [= Asia Minor] (probably ex. coll. J. Carl). ZMUC: M. adanense, 4 ♂♂, Turkey, vil. Antalya, Irmasàn Geçidi, 1430–1520 m, 28.IV.1982, A. Vigna Taglianti leg., H. Enghoff det. Diagnosis. Differs from M. serratum and M. turcicum by the smooth, rather than serrated, anterior opisthomere process; from M. tenenbaumi and M. arcuatum sp. n. by the apically gradually convex, rather than incised, promere; and from M. genezarethanum by the posterior opisthomere process (pp) being subequal to, rather than shorter than, solenomere (s), and by the smooth anterior solenomere protrusion (asp) devoid of plumose outgrowths. Redescription. Measurements: syntype (MNKB) ♂ in stadium XI, body ring number 58+1+T, body length 27 mm, height 2.1 mm; syntype ♀ (MNKB) in stadium XI, body ring number 57+1+T, body length 35 mm, height 2.3 mm; probable syntype ♂ (NHMW) in stadium XII, body ring number 53+1+T, length 35.1 mm, height 2.1 mm; ♀ (NHMW) in stadium XII, body ring number 51+1+T, length 35.6 mm, height 2.6 mm; non-type ♂♂ (ZMUC) in stadia X–XII, body ring number (47+3+T)–(55+1+T), length 26.7–40.8 mm, height 1.7–2.2 mm. Coloration: Somewhat faded, overall brownish, with a narrow, blackish, mid-dorsal stripe and broad, short, dark, transverse stripes on prozona extending from ozopore level up to some distance from mid-dorsal stripe. External structures: 20 labral setae. Gnathochilarium typically julid, promentum relatively large, nearly 2 times longer than broad. Collum smooth, with 3–4 short scratches near posterolateral corner. Prozona with numerous short, parallel scratches; metazona densely striated, with 10 striae in a square with sides equal to metazonal length just below ozopore level; posterior margin with one moderately dense row (ca 2/3 to equal to metazonal length) of setae. Ozopores placed just behind sutures between pro- and metazona, suture itself being straight in anterior body rings, slightly sinuous in posterior ones. Tarsi of midbody legs slightly shorter than tibiae. Anal valves with moderately dense, very long pilosity; no distinct rows of short setae along anal valves’ caudal margins. Pre-anal process long, straight, shorter and stouter in females; with a long, thin, somewhat down-turned, hyaline tip, slightly surpassed by the longest setae on anal valves. Subanal scale triangular, pointed, strongly protruding caudally behind anal valves’ contour by almost half of its length in males (Fig. 1); much shorter and blunter in females. Male sexual characters: Mandibular stipites enlarged, protruding mostly anteriad, forming a distinct rounded anterior corner. Leg-pair 1 typical, parallel hooks with somewhat diverging tips; tarsal remnants absent. Pleurotergum 7 ventrally with rather short, rounded lobes at border between pro- and metazona, protruding mostly mesad behind opisthomeres (Figs 2–3). Walking legs with pads on femur, postfemur and tibia; postfemoral and tibial ones undulate, femoral one small, flattened. Gonopods (Figs 2, 4–7): Promere (Fig. 5 and P in Figs 2 and 4) more than 3 times higher than broad, gradually narrowing distally, with a broadly rounded tip; a prominent median ridge (r) and a deep, narrow, median groove (g) on caudal face. Flagellum (fl in Fig. 5) slightly longer than promere, thin in distal half. Opisthomere (Figs 4, 6–7) strongly elongated, basally curved posteriad, bearing two processes: a large, flat posterior process (pp) subequal in length to solenomere (s) and bearing several blunt teeth at its apical margin; and a spiniform anterior process (ap) deviating mesad. Solenomere simple, tubular, with a short anterior protrusion (asp). Several spine-like filaments (sf) present basally along flagellum channel (fc), these looking more setiform apically. Gonopods in situ (Fig. 2): conspicuously protruding outside gonopod sinus, their tips reaching up to coxae 11 or 12; promere slightly outreaching solenomere; both promeres tightly connected to one another by transverse muscles and by a small chitinous knot between their bases, promere-opisthomere connection weaker. Female sexual characters: First two leg-pairs slightly enlarged. Vulvae (Fig. 8) about as high as broad, cylindrical, somewhat tapering apically; bursa almost completely symmetrical, with a distinct apical margin, a soft median groove and an ellipsoid opening (o) not exceeding the margin; operculum (op) subequal to bursa, its


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FIGURES 1–9. Megaphyllum curvifolii males (Figs 1–7; 9) from Irmasàn Geçidi, Turkey and syntype female (Fig. 8) from “Cilicien”, south-central Turkey: (1) telson; (2) pleurotergum 7 and gonopods in situ, lateral view; (3) right pleurotergal protrusion, ventral view; (4) right gonopods, postero-mesal view; (5) connected promeres, posterior view; (6) distal half of left opisthomere, mesal view; (7) left opisthomere, antero-apical view; (8) left vulva, mesal, slightly anterior view (receptaculum seen by transparency, but drawn with solid lines for better emphasis); (9) penis, posterior view. Abbreviations: ap: anterior process, asp: anterior protrusion of the solenomere, at: apodematic tube, bl: bursal lamellar protrusions, ct: connecting tube, fc: flagellum channel, fl: flagellum, g: median groove, o: opening, op: operculum, opl: operculum’s lamellar protrusions, P: promere, pa: posterior ampulla, ph: posterior hump, pp: posterior process, r: median ridge, s: solenomere, sf: spine-like and setiform filaments. Scale bars of the drawings: 0.2 mm. REDESCRIPTION OF SUBGENUS PARANCISTRUM


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hyaline protrusions (opl) barely exceeding bursal ones (bl). Inner structures of vulva consisting of a large, globular, posterior ampulla (pa) joining a finger-shaped apodematic tube (at) through a thin, spirally folded, connecting tube (ct). Distribution. Turkey: Cilicien (type locality of M. curvifolii) (Verhoeff 1898) [between Mersin and Tarsus (cf. Hoffman & Lohmander1964)]; Adana, near the bay of Iskenderun (type locality of M. adanense) (Verhoeff 1943); vil. Mersin, between Tarsus and Gülek, environs of Tasobagi; Mersin, Taurus, environs of Çamalan; vil. Mersin, environs of Çamliyayla; vil. Konya, environs of Mus; vil. Antalya, Irmasan Geçidi; vil. Antalya, Topraktepe; vil. Antalya, Korküteli (localities for M. adanense) (Enghoff 2006). Remarks. Studying the type material leaves no doubt that Verhoeff failed to recognize M. curvifolii when he, 45 years later, described M. adanense. Penes were only investigated in the non-type males. The only successfully dissected and drawn penis (Fig. 9) was very soft, transparent and apparently hollow. It was elongated, straight, with two short parallel lobes and two short triangular lamellae—one was probably broken off during dissection. This rather unusual (compared to other examined species of the tribe Brachyiulini) shape may be the result either of a malformation or a normal alteration concerning the period immediately before or after copulation. Another dissected male seemed to have a very similar penis to the one observed in M. turcicum (Figs 35 and 36), but it was accidentally lost before being depicted. However, we cannot say anything definitive regarding the penis of M. curvifolii pending further investigations.

Megaphyllum (Parancistrum) genezarethanum (Verhoeff, 1923) comb. nov. Figs 10–18 and 57 Brachyiulus genezarethanus Verhoeff, 1923: 131–132. Chromatoiulus genezarethanus: Jawłowski 1931: 165. Chromatoiulus palaestinus Jawłowski, 1931: 164–165, figs 11–13, syn. n. Chromatoiulus (Phauloiulus) palestinus (sic!): Attems 1940: 307. Megaphyllum palaestinus (sic!): Tăbăcaru 1995: 25. Brachyiulus (Chromatoiulus) genezarethanus: Tăbăcaru 1995: 26.

Material examined. ZSM: genezarethanum 1 ♀ holotype (missing caudal part), Reg. Nr. A20060676, Asia, Israel, Lake Genezareth [= Lake of Galilee]; ZMUC: palaestinum 1 ♂, 1 ♀, Israel, Allonim, Lower Galil, 29.X.1988, M. R. Warburg leg. & ded., H. Enghoff det. ZMTA: M. genezarethanum, 1 ♂, Elyaquim, 5.II.1990, leg. ?, BV det. Diagnosis. Differs from M. serratum and M. turcicum by the smooth, rather than serrated, anterior opisthomere process; from M. tenenbaumi and M. arcuatum sp. n. by the apically narrowly rounded, rather than incised, promere; and from M. curvifolii by the posterior opisthomere process (pp) being shorter than, rather than subequal to, solenomere (s), and by the anterior solenomere protrusion (asp) being not smooth, but bearing plumose outgrowths. Descriptive notes. The examined specimens are somewhat longer and thicker compared to the measurements given by Verhoeff (1923) and Jawłowski (1931): male in stadium X, 57+1+T, body length 42 mm, height 2.4 mm; female from the ZMUC in stadium XII, 52+1+T, body length 38 mm, height 3 mm. 24 labral setae. Male mandibular stipites considerably enlarged (Fig. 10), broadly rounded, without distinct anterior corner. Metazona with usual marginal whorl of setae, in disagreement with Verhoeff’s (1923) observation of “almost completely bare edges of body rings”. Penis stout, somewhat longer than broad, with short, parallel, apical lobes and quite long, blunt, terminal lamellae directed somewhat caudad (Figs 12–13). Gonopods: Promere (Fig. 15; P in Fig. 14) strongly elongated, more than 3 times longer than broad, gradually narrowing towards apex, the latter narrowly rounded. Flagellum subequal in length to promere. Opisthomere (Figs 14 and 16) with a flattened, slightly anteriad bent, posterior process (pp) ending up by a rounded, gently striated apex, and a smooth, thin, pointed, anterior process (ap) reaching just below solenomere (s). Solenomere slightly bulbous, with a lamellar anterior protrusion (asp) ending up by several plumose outgrowths. Pro- and opisthomere subequal in situ.


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FIGURES 10–18. Megaphyllum genezarethanum male (10–16) and female (17) from Allonim, Israel, and holotype female (18) from Lake of Galilee, Israel: (10) head with anterior body rings (antennae broken; not to scale); (11) telson with last body rings (not to scale); (12) and (13) penis, lateral and posterior views, respectively; (14) distal half of left gonopods, mesal view; (15) right promere, posterior view; (16) right opisthomere, mesal view; (17) right vulva, lateroposterior view; (18) receptaculum of left vulva. Abbreviations: ap: anterior process, asp: anterior protrusion of the solenomere, at: apodematic tube, bl: bursal lamellar protrusions, ch: central hump, ct: connecting tube, fc: flagellum channel, fl: flagellum, g: median groove, lh: lateral hump, mh: mesal hump, o: opening, opl: operculum’s lamellar protrusions, P: promere, pa: posterior ampulla, ph: posterior hump, pp: posterior process, r: median ridge, s: solenomere. Scale bars of the drawings: 0.2 mm.

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Vulvae (Figs 17 and 57) densely setose, cylindrical, almost completely symmetric, lateral hump (lh) slightly higher than mesal one (mh). Opening (o) small, slit-like, placed just on top, not reaching apical margin. Bursal apical margin nearly straight, bursal side-lobes protruding into very large, rounded, hyaline lamellae (bl). Operculum straight, slightly shorter than bursa, protruding into a bifurcated hyaline lamella (opl), somewhat shorter than bursal ones. In situ operculum directed mostly anteriad and less strongly laterad. Receptaculum seminis (Fig. 18) consisting of a wide apodematic tube (at) with a somewhat tapering bottom, and a sac-like globular posterior ampulla (pa) with a small additional finger-like reservoir, joining at through a very thin, spirally folded, connecting tube (ct). Distribution. Israel: Lake Genezareth [= Lake of Galilee] (type locality of M. genezarethanum) (Verhoeff 1923); Haifa, Mt. Karmel (type locality of M. palaestinum) (Jawłowski 1931); Allonim, Elyaquim (new data). Remarks. Although a synonymy based on female characters is always risky, in this case the vulvae of the M. genezarethanum holotype proved to be virtually identical to those of the female from ZMUC, and clearly differing from the examined structures in all other consubgeners.

Megaphyllum (Parancistrum) serratum (Verhoeff, 1898) Figs 19–22 Brachyiulus serratus Verhoeff, 1898: 297–298, fig. 7. Brachyiulus serratus: Verhoeff 1923: 131. Chromatoiulus serratus: Attems 1927: 229–230, figs 301–302. Chromatoiulus (Phauloiulus) serratus: Attems 1940: 307. Megaphyllum serratum: Enghoff 2006: 183.

Material examined. MNKB: Cat. No. 2265, syntypes 1 ♂, 1 ♀, 1 juv.; Verhoeff syntype slide No. 1201: gonopods + leg-pair 2. Diagnosis. Differs from M. curvifolii, M. genezarethanum, M. tenenbaumi, and M. arcuatum sp. n. by the serrated, rather than smooth, anterior opisthomere process; and from M. turcicum by the narrow posterior opisthomere process which is slightly shorter than solenomere, as opposed to the same process being very wide, higher than solenomere in the latter species. Descriptive notes. According to Verhoeff’s (1898) original description, this species seems to be comparable in size with M. transsylvanicum. However, the examined specimens proved to be considerably smaller: male in developmental stadium XII, 52+1+T, body length 29 mm, height 1.7 mm; female in stadium XII–XIII, 48+1+T, length 33 mm, height 2.2 mm. 20 labral setae. Collum with 4–5 shallow striae near posterolateral corner; male mandibular stipites fairly large, protruding mostly ventrad, with a rounded infero-anterior corner. Ozopores placed just behind suture between pro- and metazona, slightly sinuous in anterior body rings; male pleurotergum 7 almost unmodified, only its ventral metazonal edge slightly convex. Metazona deeply striated, with 10 striae in a square with sides equal to metazonal length just below ozopore level; metazonal hind margins densely pilose. Pre-anal process directed slightly ventrad, rather short, strongly robust, surpassed by longest setae on anal valves. Subanal scale triangular, pointed, protruding by ca 1/3 of its length behind anal valves’ rear contour in the male, blunt, not protruding in the female. Anal valves densely setose, with a distinct row of short, stiff setae along caudal margin of each valve. Gonopods: Promere (Fig. 20 and P in Fig. 19) ca 2 times longer than broad, with a rounded incision at distolateral margin; median groove (g) very deep and narrow, tightly subtending opisthomere in situ. Flagellum considerably longer than promere. Opisthomere (Figs 19, 21) with a narrow, slightly tapering, posterior process (pp), with several blunt teeth at distomesal margin; a flattened, mesad deviating, anterior process (ap) with a serrated apical part; and a slender, tubular solenomere (s) with a finger-like anterior protrusion (asp). Promere in situ considerably shorter than opisthomere; promere-opisthomere connection exceptionally strong due to a chitinous knot at their bases, similar to those normally observed between promeres of most julids. Vulvae (Fig. 22) strongly asymmetric: lateral hump of bursa (lh) protruding apically high above both mesal hump (mh) and operculum. Short, rounded hyaline lamellae present apically on both bursa and operculum. Setation dense all over vulva. Shape of receptaculum (at, pa) typical of the subgenus, but smaller in relation to vulva size compared to condition observed in the other species.


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FIGURES 19–22. Megaphyllum serratum syntypes, male (19–21) and female (22): (19) right gonopods, mesal, slightly anterior view; (20) right promere, posterior view; (21) right opisthomere, meso-anterior view; (22) right vulva, lateroposterior view (setae omitted, receptaculum seen by transparency, but drawn with solid lines for better emphasis). Abbreviations: ap: anterior process, asp: anterior protrusion of the solenomere, at: apodematic tube, ch: central hump, ct: connecting tube, fc: flagellum channel, fl: flagellum, g: median groove, lh: lateral hump, mh: mesal hump, P: promere, pa: posterior ampulla, ph: posterior hump, pp: posterior process, r: median ridge, s: solenomere, sf: spine-like filaments. Scale bars: 0.2 mm.



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Distribution. Turkey, Pernata, Inevi (type locality) (Verhoeff 1898; Enghoff 2006). Remarks. At the present the examined individuals are all strongly faded, but, according to Verhoeff (1898), only the males have yellow-reddish dorsolateral sides which might suggest sexual dimorphism.

Megaphyllum (Parancistrum) tenenbaumi (Jawłowski, 1931) Figs 23–34 and 56 Chromatoiulus tenenbaumi Jawłowski, 1931: 163–164, figs 8–10. Chromatoiulus (Phauloiulus) tenenbaumi: Attems 1940: 307. Megaphyllum tenenbaumi: Tăbăcaru 1995: 25.

Material examined. Neotype (ZMTA) (Figs 23–25): 1 ♂ (intact), Israel, Nahal Keziv [Nahal Kziv], pitfall trapping, 134-5, 17– 31.I.1999, leg. M. Finkel., SG det. Additional material. All from Israel: ZMUC: 1 ♀, unknown locality, probably M. R. Warburg leg., SG det. ZMTA: M. Finkel leg., SG det.: 1 ♂, 3 ♀♀, Golan; from Nahal Kziv: 3 ♂♂, 1 juv., pitfall trapping, SF33-3, 17– 31.I.1999; 2 ♀♀, NFS7-2,17–31.I.1999; 1 ♀, pitfall trapping, 134-4, 17–31.I.1999; 1 ♂, SFS1-7, 12.XII.1998– 2.I.1999; 1 ♂, NFS6-1, 17–31.I.1999; 1 ♀, pitfall trapping, NFS9-9, 17–31.I.1999; 1 ♂, pitfall trapping, SFS1-10, 12.XII.1998–2.I.1999; 1 ♀, 1 juv., pitfall trapping, NFS6-7, 12.XII.1998–2.I.1999;1 ♀, 1 juv., pitfall trapping, NFS6-9, 17–31.I.1999; 1 ♀, pitfall trapping, NSF7-4, 12.XII.1998–2.I.1999; 1 ♀, pitfall trapping, B4-2, 12.XII.1998–2.I.1999; 1 ♂, 3 ♀♀, 1 juv., pitfall trapping, NFS5-6, 12.XII.1998–2.I.1999; 3 ♀♀, pitfall trapping, NFS5-9,17–31.I.1999; 2 ♂♂, 1 ♀, 2 juv., pitfall trapping, NFS7-2, 12.XII.1998–2.I.1999; 1 ♂, SFS3-1, 17– 31.I.1999; 2 ♀♀, NFS6-8, 17–31.I.1999; 1 ♀, B4-1, 12.XII.1998–2.I.1999; all M. Finkel leg.; ZMUM: 1 ♂, Nahal Kziv, pitfall trapping, B-7, 28.XI.–12.XII.1998, M. Finkel leg., SG det.; NMNHS: 1 ♂, 1 ♀, Nahal Kziv, pitfall trapping, SFS2-10, 12.XII.1998–2.I.1999, M. Finkel leg., SG det.; HNHM: 1 ♂, Nahal Kziv, pitfall trapping, SFS3-7, 17–31.I.1999, M. Finkel leg., SG det. Diagnosis. Differs from M. serratum and M. turcicum by the smooth, rather than serrated, anterior opisthomere process; from M. curvifolii and M. genezarethanum by the apically incised, rather than rounded, promere; and from M. arcuatum sp. n. by the anterior solenomere protrusion (asp) being short, blunt and pointing distad, as opposed to the same process being slenderer, tapering, turned posteriad in the latter species, as well as by the promere’s incision being considerably deeper than in M. arcuatum. Descriptive notes. Measurements similar to those observed in M. genezarethanum: males, 33–40 mm in length, 2.1–2.4 mm in height, females, 35–45 mm, and 2.9–3.2 mm, respectively. The two species are also alike in coloration and external characters (Figs 23–24); pre-anal process and subanal scale somewhat longer, more strongly protruding in M. tenenbaumi (Fig. 25). Penis (Figs 26–27) with more strongly converging—rather than parallel—sides and with almost entirely caudad directed lamellae. Gonopods: Promere (Fig. 29 and P in Fig. 28) somewhat stouter, less strongly elongated than in genezarethanum, with a deeply incised apex. Opisthomere (Figs 28 and 30) with a slightly tapering posterior process (pp) ending up by a narrowly rounded apex; a pointed anterior process (ap) slightly broader and longer (almost reaching the level of solenomere’s tip) than that in genezarethanum. Solenomere (s) with a short, anterior, lamellar protrusion (asp), without apical plumose outgrowths. Vulvae (Figs 31–34 and 55) stouter, more robust, with more prominent mesal (mh) and lateral apical humps (lh), and shorter bursal lamellae (bl). The scanned vulva of one female showed a bizarre modification of part of its setae: bulging, twisted-like knots in their mid-sections (Fig. 33). However, the vulvae of the other examined female were perfectly normal, so these modified setae are most likely an aberration rather than a species-specific character. Inner structure of vulvae as in Fig. 34; main differences from M. genezarethanum being the distally wider apodematic tube (at), the more globular posterior ampulla (pa) and the thicker connecting tube (ct). Distribution. Israel: Rosh-Pina [Rosh Pinna] (type locality) (Jawłowski 1931); Golan Heights, Nahal Kziv (new records). Remarks. The type specimens of M. tenenbaumi had been deposited, like most of the other type material of Jawłowski, in the Krakow Museum, but everything perished during World War II. (J. Wytwer, personal communications). As there are two particularly similar species—M. genezarethanum and M. arcuatum sp. n.—a name-bearing type is necessary to clarify the taxonomic status of M. tenenbaumi. This situation requires a neotype


VAGALINSKI ET AL.

FIGURES 23–34. Megaphyllum tenenbaumi neotype male (23–25), and non-type male (26–30) from Nahal Kziv, Israel, and female from ZMUC (31–33), unknown locality: (23) habitus; (24) head with anterior body rings; (25) telson with last two body rings; (26) and (27) penis, lateral and posterior view, respectively; (28) left gonopods, mesal view; (29) right promere, posterior views; (30) distal part of right opisthomere, lateral view; (31) right vulva, apical view; (32) right vulva, postero-lateral view; (33) altered setae on distal part of bursa; (34) receptaculum of left vulva. Abbreviations: ap: anterior process, asp: anterior protrusion of the solenomere, bl: bursal lamellar protrusions, ch: central hump, fc: flagellum channel, fl: flagellum, g: median groove, lh: lateral hump, mh: mesal hump, o: opening, op: operculum, opl: operculum’s lamellar protrusions, P: promere, ph: posterior hump, pp: posterior process, r: median ridge, s: solenomere. Scale bars of the drawings: 0.2 mm.



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designation under Article 75.1. (ICZN 2000). The neotype is chosen from Nahal Kziv, Israel, only ca 45 km away from the original type locality (Jawłowski 1931). The morphometric and descriptive characters of the neotype are as follows: Developmental stadium XI, body ring number 50+1+T, length 36 mm, height 2.1 mm. Coloration somewhat faded, antennae dark brown, head lighter brownish with a black band between ocellaria; collum brown with darker margins; prozona darker in anterior rings, greyish above, brownish below ozopore level; metazona brown, darker at ozopore level and above it at posterior margins; legs brownish, with darker distal 3 podomeres; pre-anal ring blackish-brown with a light brown caudal process; anal valves irregularly brownish proximally, yellowish-beige distally. Mandibular stipites enlarged (arrow in Fig. 24), ellipsoid, without distinct anterior corner. Metazona densely striated, with 10–11 striae in a square with sides equal to metazonal length just below ozopore level; a row of erect setae ca 1/2–2/3 of metazonal length present at metazonal hind margin. Pre-anal process straight (Fig. 25), with a slightly ventrad turned hyaline tip surpassing the longest setae on anal valves. Anal valves densely pilose, without distinct rows of shorter setae along caudal margin. Subanal scale triangular, sharply pointed, protruding by about its distal quarter behind rear contour of anal valves.

Megaphyllum (Parancistrum) turcicum (Verhoeff, 1898) Figs 35–40 Brachyiulus turcicus Verhoeff, 1898: 301, fig. 14. Brachyiulus turcicus: Verhoeff 1923: 131. Chromatoiulus turcicus: Attems 1927: 230. Chromatoiulus (Phauloiulus) turcicus: Attems 1940: 307. Megaphyllum turcicum: Enghoff 2006: 184.

Material examined. MNKB: ZMB 12379, syntype ♂ gonopods + 1st and 2nd leg-pairs as slide preparation, Cilicien. MHNG: 2 ♂♂, 9 ♀♀, 3 juv., Turkey, Mersin, road to Arslanköy, 5 km SE Aladag, Kerimler, 680 m, 36°54’45” N, 34°31’44” E, 10.V.2004, C. Besuchet leg., BV det. Diagnosis. Differs from M. curvifolii, M. genezarethanum, M. tenenbaumi, and M. arcuatum sp. n. by the serrated, rather than smooth, anterior opisthomere process; and from M. serratum by the very broad posterior opisthomere process which is higher than the solenomere, as opposed to the same process being significantly narrower, slightly shorter than the solenomere in the latter species. Descriptive notes. Measurements: males in developmental stadia X–XI, body ring number (52-53+2+T), body length 27–29 mm, height 1.6–1.7 mm; females in developmental stadia XI with 2 apodous body rings, 26–31 mm long, 2.1–2.3 mm high. 18 labral setae. Collum with 5–6 barely visible scratches near anterolateral corner. Metazonal setation rather short (ca. ½ or less of metazonal length), erect, not very dense. Pre-anal process very long and straight, exceeding the longest setae on anal valves; not roof-like, but narrow, with a sharp ventral keel. Subanal scale triangular, pointed, protruding by ca 1/3 of its length behind caudal contour of anal valves in males, blunt and barely protruding in females. Anal valves with moderately dense pilosity on surface and a distinct row of shorter setae along caudal margin of each valve. Male mandibular stipites protruding more anteriad than ventrad, forming a distinct, rounded, anterior corner. Male pleurotergum 7 with weakly protruding, broadly rounded lobes lateral to gonopods. Penis (Figs 35–36) somewhat longer than broad, with slightly converging sides; anterocaudally constricted just before apical lobes, the latter short, parallel, blunt pyramidal; terminal lamellae thin, rounded, directed entirely dorsad. Promereopisthomere connection stronger than promere-promere one. Gonopods: Promere (Fig. 38, P in Fig. 37) ca 2.5 times longer than broad, slightly and gradually narrowing towards apex, the latter widely rounded; median ridge (r) and median groove (g) well-pronounced. Flagellum (fl) considerably longer than promere. Opisthomere (Figs 37, 39) with a very large, shield-like, posterior process (pp), postero-mesally covered with short and stout spines; a slender, s-bent anterior process (ap) with a serrated distal part. Solenomere (s) turned distad, bearing a bifurcated, pointed, anterior protrusion (asp). Gonopods in situ: promere subequal to solenomere, both surpassed by posterior process; promere-opisthomere connection stronger than promere-promere one.


VAGALINSKI ET AL.

FIGURES 35–40. Megaphyllum turcicum male (35–39) and female (40) from near Aladağ, Turkey: (35) and (36) penis, caudal and lateral views, respectively; (37) right gonopods, mesal view; (38) left promere, posterior view; (39) left opisthomere, anterior, slightly mesal view; (40) right vulva, posterior, slightly mesal view (receptaculum seen by transparency, but drawn with solid lines for better emphasis). Abbreviations: ap: anterior process, asp: anterior protrusion of the solenomere, at: apodematic tube, bl: bursal lamellar protrusions, ct: connecting tube, fc: flagellum channel, fl: flagellum, g: median groove, lh: lateral hump, mh: mesal hump, opl: operculum’s lamellar protrusions, P: promere, pa: posterior ampulla, ph: posterior hump, pp: posterior process, r: median ridge, s: solenomere, sf: spine-like filaments. Scale bars: 0.2 mm.

Vulvae (Fig. 40) subcylindrical, compact, slightly higher than maximum width, mostly symmetric; lateral valve somewhat broader than mesal one. Bursa gently constricted before its distal 1/3, apical margin with weakly pronounced lateral and mesal humps (lh and mh); central hump absent. Bursal sidelobes ending with short,



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rounded, hyaline protrusions. Opening small and ellipsoid. Operculum subequal in height to bursa, without hyaline protrusions; setation on operculum denser than on bursa. Receptaculum seminis consisting of a rather narrow apodematic tube (at), a large, sac-like posterior ampulla (pa) joining the apodematic tube through a strongly folded, but not spirally twisted connecting tube (ct). Distribution. Cilicien [between Mersin and Tarsus (cf. Hoffman & Lohmander 1964)] (type locality) (Verhoeff 1898). Remarks. This is the second record of the species since its original description, again from the terra typica.

Megaphyllum (Parancistrum) arcuatum sp. n. Figs 41–55 Holotype. (ZMTA) (Figs 41–43): 1 ♂ (intact), Adolam Nature Reserve, nr. Beit Gurvin, pitfall trapping, No. 16(8), XI.2001, Y. Mandelik leg., SG det. Paratypes. (ZMUC): 2 ♂♂ (one intact, the other in head-collum+ first 2 body rings +3rd– 6th body rings + rest of body, gonopods dissected, 1 half prepared for SEM), 2 ♀♀ (one unbroken, the other fragmented, with dissected vulvae), 2 juv. ♂♂, Israel, Segev, X.1994, leg. M. Warburg; (ZMUM): 1 ♂ (intact), Israel, Sdot Micha, pitfall traps, No. 45(12), XI.2001, leg. Y. Mandelik; (NMNHS): 1 ♂ (unbroken), Israel, Adolam Nature Reserve, nr. Beit Gurvin, pitfall trapping, No. 18(7), XI.2001, Y. Mandelik leg., SG det.; (HNHM): 1 ♂ (intact), Israel, Adolam Nature Reserve, nr. Beit Gurvin, pitfall trapping, No. 17(4), XI.2001, Y. Mandelik leg., SG det.; (ZMTA): 1 ♀ (intact), Israel, Adolam Nature Reserve, nr. Beit Gurvin, pitfall trapping No. 16(3), XI.2001, Y. Mandelik leg., SG det. Other material. All from Israel, pitfall trapping: ZMTA: From Sdot Micha: 1 ♂, No. 42 A(6), XI.2001, Y. Mandelik leg., SG det.; 1 ♂, 2 ♀♀, No. 45(7), Y. Mandelik leg., SG det. From Ramot Avishur, nr. Beit Gurvin: 1 ♂, No. 24(8), XI.2001, Y. Mandelik leg., SG det.;1 ♂, No. 28(10), XI.2001, Y. Mandelik leg., SG det.; Adolam Nature Reserve, nr. Beit Gurvin: 1 ♂, No. 4(9), road; 1 ♂, No. 16(9), road; 1 ♂, No. 6(10), road; 1 ♀, No. 14(10); 1 ♀, No. 4(6). ZMUC: 1 ♂, Segev, Lower Galil, 1994, M. Warburg leg., H. Enghoff det. as cf. tenenbaumi; 1 ♂, 1 ♀, Segev, Lower Galil, 29.I.1986, M. R. Warburg leg. & ded., H. Enghoff det. as cf. tenenbaumi. Diagnosis. Differs from M. serratum and M. turcicum by the smooth, rather than serrated, anterior opisthomere process; from M. curvifolii and M. genezarethanum by the apically incised, rather than rounded, promere; and from M. tenenbaumi by the anterior solenomere protrusion being slenderer, tapering and turned posteriad, as opposed to the same process being short, blunt and pointing distad in the latter species, as well as by the promere’s apical incision being considerably more shallow than in M. tenenbaumi. Name. To emphasize not only the fact that the entire gonopod is clearly bent caudad, like in the above two species plus M. curvifolii, but also the arcuate anterior solenomere protrusion (asp), which is unique in the subgenus. Description. Measurements: holotype ♂ in stadium XI, 53+1+T, length 37 mm, height 2.1 mm. Paratype ♂♂ (ZMUC): in stadium XI, 53+1+T, length 34 and 31 mm, height 2.1 and 2.2 mm respectively; (ZMUM): in stadium XI, 52+2+T, length 30 mm, height 1.9 mm; (NMNHS): in stadium XI, 54+1+T, length 42 mm, height 2.2 mm; (HNHM): in stadium XI, 51+1+T, length 35 mm, height 2.1 mm. Paratype ♀♀ (ZMUC): in stadium XII, 55+1+T, length 37 mm, height 2.9 mm and 3.1 mm; (ZMTA): in stadium XI, 51+1+T, length 31 mm, height 2.1 mm. Coloration: Overall brownish (Fig. 41); head (Fig. 42) dark brown with a black band between ocellaria; collum dark brown with blackish margins; metazona castaneous brown with ochre-brown hind margins; prozona irregularly striped light and dark brown; a dark grey, continuous, mid-dorsal stripe divided by a black axial line, and two blackish, broken, dorsolateral stripes at ozopore level. External structures: 2 frontal, 4 supralabral and 22 labral setae. Antennomeres 5, 4 and 3 subequal in length. Gnathochilarium typically julid, with 3 apical setae on each stipes and 4–5 setae on each lingual plate; promentum relatively large, length to width ratio: 1.9–2.1. Collum smooth, with only one long stria running parallel to anterolateral margin. Prozona with numerous short, parallel scratches; metazona densely striated, with 10–11 striae in a square with sides equal to metazonal length just below ozopore level; a moderately dense row of erect setae ca 1/2–2/3 of metazonal length at metazonal hind margin. Ozopores placed just behind suture between pro- and metazona, suture slightly sinuous in front of them around midbody and further behind. Tarsi of midbody legs


VAGALINSKI ET AL.

FIGURES 41–45. Megaphyllum arcuatum holotype male (40–42) and paratype female from Segev (43–44): (41) habitus (not to scale); (42) head with anterior body rings; (43) telson with last three body rings; (44) left vulva, posterior view; (45) right vulva, lateral, slightly posterior view (setae omitted, receptaculum seen by transparency, but drawn with solid lines for better emphasis). Abbreviations: at: apodematic tube, bl: bursal lamellar protrusions, ch: central hump, ct: connecting tube lh: lateral hump, mh: mesal hump, o: opening, op: operculum, opl: operculum’s lamellar protrusions, pa: posterior ampulla, ph: posterior hump. Scale bars: 0.3 mm (Fig. 44) and 0.2 mm (Fig. 45).



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slightly shorter than to equal to tibiae. Pre-anal process long, straight, surpassing the longest setae on anal valves, with a well-developed ventral keel and a distinct, thin, pointed, hyaline tip turned slightly ventrad (broken off in most specimens). Subanal scale triangular, pointed, protruding by about its distal quarter behind rear contour of anal valves in males, blunt, not protruding in females. Anal valves moderately densely pilose, without distinct rows of shorter setae along caudal margin (Fig. 43).

FIGURES 46–54. Megaphyllum arcuatum paratype male (46–47; 49–40; 52–54) from Segev and non-type male (48, 51) from near Beit Gurvin: (46) and (47) penis, caudal and lateral views, respectively; (48) right gonopods, mesal views; (49) joint promeres, posterior view; (50) left promere, posterior view (51) left opisthomere, lateral view; (52) distal part of right opisthomere, mesal view; (53) distal part of right opisthomere, postero-mesal view; (54) distal part of right opisthomere, anterior view. Abbreviations: ap: anterior process, asp: anterior protrusion of the solenomere, c: neck-like constriction, fc: flagellum channel, fl: flagellum, g: median groove, P: promere, pp: posterior process, r: median ridge, s: solenomere, sf: spine-like filaments. Scale bars of the drawings: 0.2 mm.


VAGALINSKI ET AL.

FIGURES 55–57. Schematic bursal contours of left vulvae in posterior view of (55) M. arcuatum, (56) M. tenebaumi and (57) M. genezarethanum. Abbreviations: ch: central hump, lh: lateral hump, mh: mesal hump. Scale bar: 0.2 mm.

Male sexual characters: Mandibular stipites enlarged (arrow in Fig. 42), ellipsoid or egg-shaped, equally protruding ventrad and anteriad, with a regularly convex ventral margin and a straight to slightly concave anterior margin, forming no distinct anterior corner. Leg-pair 1 normal parallel hooks with somewhat diverging tibial outgrowths; tarsal remnants not evident. Leg-pair 2 with two simple adhesive pads, one each on postfemur and tibia. Pleurotergum 7 ventrally with large, trowel-shaped lobes protruding entirely mesad and closing gonopod sinus from behind. Walking legs with pads on femur, postfemur and tibia; femoral pads strongly flattened, gradually diminishing in size towards telson. Penis (Figs 46–47) longer than broad, with parallel, tightly contiguous, somewhat posteriad turned, apical lobes about half as long as penis, ending up with small, rounded, terminal lamellae. Gonopods: Promere (Figs 49–50 and P in Fig. 48) ca 3 times longer than broad, more or less regularly wide until a laterally sloping, gently incised apex; median ridge (r) prominent, robust, rather short; median groove (g) broad and rather deep. Flagellum (fl in Figs 48–49) somewhat longer than promere, thin. Opisthomere (Figs 48, 52–54) slender, basally somewhat curved posteriad, with a well-detached, parallel, posterior process (pp) whose distal part is slightly widened, ending up in a thick, rounded margin and bearing several small humps; and a slender, tapering, somewhat dorsoventrally compressed, anterior process (ap). Solenomere (s) well-differentiated from main opisthomere body through a neck-like constriction (c in Fig. 52), bulging anteriorly, ending up by an anterior lamellar protrusion (asp) with a pointed, posteriad curved tip. Spine-like filaments (sf) along flagellum channel (fc) present. Gonopods in situ: largely external, protruding posteriad to reach coxae 12 (Fig. 42); solenomere slightly overreaching promere (Fig. 48). Promere-promere connection stronger than promereopisthomere one, promeres fused basally at a nearly right angle (Fig. 49), partly covering opisthomeres laterally. Female sexual characters: Leg-pair 2 and, especially, pair 1 of mature females somewhat enlarged. Vulvae (Figs 44–45, 55) cylindrical, almost completely symmetric, lateral valve of bursa distally protruding into a more prominent hump (lh) than mesal (mh) one. Bursa forming a distinct, subrectangular, apical margin. Opening (o) small, slit-like, placed just on top of bursa; bursal sidelobes ending by large, rounded, hyaline protrusions (bl). Operculum (op) narrow, subequal in height to bursa, ending up by a bifurcated hyaline protrusion. Receptaculum seminis double, consisting of a wide apodematic tube (at) ending with a short, finger-like reservoir and a globular posterior ampulla (pa) joining at just before the opening through a thin, spirally folded, connecting tube (ct). Distribution. The species seems to be endemic to Israel. Remarks. The new species shows more similarities to M. tenenbaumi than to M. genezarethanum due to the more or less clearly incised promere apex, as well as the angled connection between both promeres and the more robust, mesolaterally less strongly compressed vulvae.

Key to males of the subgenus Parancistrum, based on gonopod structure 1 -

Anterior process (ap) serrated . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Anterior process (ap) smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3



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2 3 4 5 -

Posterior process (pp) very wide and considerably exceeding solenomere (s) (Figs 37, 39) . . . . . . . . . . . . . . . . . . M. turcicum Posterior process (pp) narrow and slightly shorter than solenomere (s) (Figs 19, 21) . . . . . . . . . . . . . . . . . . . . . . M. serratum Promere with a more or less incised apical margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Promere gradually convex apically . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Promere’s apical margin shallowly incised (Figs 48–50), anterior solenomere protrusion (asp) more slender, tapering, turned posteriad (Figs 48, 51–54) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. arcuatum sp. n. Promere’s apical margin deeply incised (Figs 28–29), anterior solenomere protrusion (asp) shorter, blunter, pointing distad (Figs 28, 30) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. tenenbaumi Posterior process (pp) subequal to solenomere(s); anterior solenomere protrusion (asp) smooth (Figs 4, 6–7) . . . M. curvifolii Posterior process (pp) shorter than solenomere (s); anterior solenomere protrusion (asp) with apical plumose outgrowths (Figs 14, 16) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. genezarethanum

Key to females of the subgenus Parancistrum, based on vulvae and external morphological characters 1 2 3 4 5 -

Vulvae markedly asymmetric, with lateral hump (lh) strongly protruding distally, by far outreaching central (ch) and mesal (mh) humps, and operculum (o) (Fig. 22) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. serratum Vulvae more or less symmetric . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Vulvae with lateral (lh) and mesal humps (mh), but without central hump at apical margin of bursa . . . . . . . . . . . . . . . . . . . . . 3 Vulvae with lateral (lh), mesal (mh) and central humps (ch) at bursal apical margin (Figs 55–57) . . . . . . . . . . . . . . . . . . . . . . 4 Bursa somewhat constricted in its distal part (Fig. 40); operculum shorter than bursa . . . . . . . . . . . . . . . . . . . . . . . M. turcicum Walls of bursa more or less straight; operculum equal in height to bursa (Fig. 8) . . . . . . . . . . . . . . . . . . . . . . . . . . . M. curvifolii Vulva more gracile, mesolaterally strongly compressed, with a visibly concave median side lobe (Fig. 57); bursal hyaline protrusions (bl) very long, ca 1/3 of base-to-apex height (Fig. 17) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. genezarethanum Vulva more robust, mesolaterally less strongly compressed, with a flat, mesal side lobe; bursal hyaline protrusions shorter than 1/3 of base-to-apex height . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Subanal scale triangular, protruding behind rear contour of anal valves; bursa of vulva more asymmetric, its lateral hump (lh) much wider than mesal one (mh) (Fig. 56) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. tenenbaumi Subanal scale narrowly rounded, not protruding behind rear contour of anal valves; bursa of vulva less strongly asymmetric, lateral hump (lh) slightly wider than mesal one (mh) (Figs 44, 54) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .M. arcuatum sp. n.

Discussion The subgenus Parancistrum as redefined here displays a coherent distribution pattern extended along the eastern Mediterranean coast, with only M. serratum found more inland in Anatolia. Given the vast gap between Turkey and Israel, it seems very likely that more species of the group will be discovered in the coastal areas of Lebanon and Syria as well. Of the six recognized consubgeners, all three Israeli species—M. genezarethanum, M. tenenbaumi and M. arcuatum sp. n.—are very similar to one another, forming a solid species group characterized by the strongly elongated, externally protruding gonopods, the tightly connected promeres, the slender, smooth, spine-like anterior process, and the rather narrow posterior process of the opisthomere. Distinct areas of occurrence are hard to outline for each of them, except perhaps for M. tenenbaumi which occurs only north of the Lake of Galilee. M. arcuatum sp. n. was mainly found in the central parts of Israel, but the single record from Segev, Lower Galilee suggests a wider distribution. The only three specified localities known for M. genezarethanum are geographically too close to outline its occurrence. However, one should keep in mind that the present-day distributions of these species must have been strongly affected by the increasing aridity and deforestation blanketing the entire Near East since prehistoric times. So what we may be witnessing now might well be only remnants of a once more widespread and diverse group. Interestingly, M. curvifolii shares more similarities with the Israeli than with the other two Turkish consubgeners (M. serratum and M. turcicum), namely, the markedly elongated gonopods and the smooth, pointed anterior process. The latter two species differ significantly by their more compact and less externally protruding gonopods, as well as by the stronger promere-opisthomere connection compared to the more easily separable promeres, and by the anterior opisthomere process being serrated. M. serratum is also remarkable on its own for its strongly asymmetric vulvae. The place of Parancistrum within the prolific genus Megaphyllum and the tribe Brachyiulini is questionable, but the nominotypical subgenus emerges as a possible sister group. The well-developed posterior opisthomere process (pp), the rather simple solenomere (s), the presence of an anterior opisthomere process (ap) (albeit not in


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all species of Megaphyllum s. str.), the markedly cylindrical vulvae with mostly apically positioned openings, as well as the overall shape of the penes, are all common traits for both groups (see also Lazányi & Vagalinski accepted). Species of Megaphyllum s. str. differ from Parancistrum (see Tab. 1) by the blunt, mostly trapezoidal subanal scale compared to its sharp triangular shape observed in the presently studied subgenus, as well as by their relatively more compact, mostly internal gonopods, with the promeres tightly enveloping the opisthomeres, the latter fitting in a deep, narrow groove on the posterior side of the promeres. However, these gonopodal characters also hold true to some extent for M. turcicum and M. serratum, which on the other hand do not show posterior solenomere process so characteristic of the subgenus Megaphyllum (Lazányi & Vagalinski accepted). Furthermore, the anterior solenomere protrusion in Parancistrum cannot be homologous to the anterior solenomere process in Megaphyllum s. str., since the process is positioned at the end of the flagellum channel, unlike the condition in Parancistrum where the channel ends up clearly behind the anterior protrusion. If the above morphological similarities prove to be apomorphic, it would be somehow difficult to explain the considerable distance between southern Turkey-Israel and the Balkans, the latter being the main distribution area of Megaphyllum s. str., with 15 of 20 species occurring there. A possible explanation might be that both subgenera represent an ancient lineage of direct descendants of the ancestors of the Brachyiulini which once might have inhabited a greater part of the Levant and Asia Minor. Species of Pontobrachyiulus Lohmander, 1939—which inevitably occupy a basalmost position among other Brachyiulini taxa in a preliminary phylogenetic analysis (BV’s unpublished results)—are also similar to Parancistrum in sharing mostly external gonopods, a massive, flattened, posterior opisthomere process and a slender, albeit simpler, solenomere. The most recent molecular studies (Enghoff et al. in press) place Megaphyllum (Donbrachyiulus) rossicum and species of the genera Brachyiulus and Acropoditius in a clade opposing the examined species of Megaphyllum s. str. This implies polyphyly of Megaphyllum, with the closest relatives of the nominotypical subgenus possibly living outside Europe. Similar studies on other members of Brachyiulini, including species of Parancistrum, would reveal further details of their phylogenetic relationships. TABLE 1. Main diagnostic differences between the subgenera Parancistrum and Megaphyllum s. str. of the genus Megaphyllum.

♂

Character subanal scale anterior process (ap) solenomere (s)

♀

vulval opening (o) posterior ampulla (pa) operculum (op)

Parancistrum triangular, pointed long, well-developed

Megaphyllum s. str. trapezoidal, with several blunt teeth apically obscure, mostly fused with main opisthomere body or fully absent with an anterior protrusion (asp) positioned with an anterior and a posterior process, the anteriad of the flagellum channel (fc) former right at the end of the flagellum channel narrow oval or slit-like, positioned entirely wider oval, usually slightly extending below apically bursal apical margin usually (except for M. arcuatum) with never with additional reservoir additional finger-like reservoir subequal to or slightly shorter than bursa significantly shorter than bursa

Acknowledgements We would like to thank Prof. Henrik Enghoff, Natural History Museum of Denmark, Zoological Museum, University of Copenhagen, Stefan Friedrich, Zoologische Staatssammlung, München, Dr. Jason Dunlop, Museum für Naturkunde, Berlin, Dr. Verena Stagl, Naturhistorisches Museum, Wien, Dr. Peter Schwendinger, Muséum d’histoire naturelle, Geneva, Prof. Eviatar Nevo, Institute of Evolution, University of Haifa, and Dr. Yael Mandelik, now the Faculty of Agriculture, Food and Environment, The Hebrew University of Jerusalem, Rehovot for their help in our access to the type and non-type specimens. Dr. Jolanta Wytwer, Museum and Institute of Zoology, Polish Academy of Sciences, Warsaw is also acknowledged for making some of the microphotographs. The visit of BV to the Zoological Museum, University of Copenhagen was supported by the European Commission’s (FP6) Integrated Infrastructure Initiative program SYNTHESYS (DK-TAF-1963).



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